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Estudios Geológicos, 67(2) julio-diciembre 2011, 179-186 ISSN: 0367-0449 doi:10.3989/egeol.40553.181

A new species of Adelpharctos (Mammalia, Carnivora, Ursidae) from the late Oligocene of the “Phosphorites du Quercy” (France) Una nueva especie de Adelpharctos (Mammalia, Carnivora, Ursidae) del Oligoceno Superior de las “Fosforitas de Quercy” (Francia)

L. de Bonis1

ABSTRACT

The genus Adelpharctos was known until now through the species A. mirus by a unique mandible (p2-m2) from the old collections of the Quercy whose geological age was unknown. New material coming from the locality of Pech-du-Fraysse completes our knowledge of the genus particularly for the maxilla and upper teeth. Adelpharctos belongs to the sub-family Hemicyoninae in the family Ursidae. It differs from the middle Miocene hemicyonines which have more massive molars and from the group Cephalo- gale-Phoberogale by some morphological characters. It seems to be a branch coming from the ancestral stem group of the sub-family. Keywords: Late Oligocene, France, Quercy, Carnivora, Ursidae, Hemicyoninae.

RESUMEN

El género Adelpharctos era solo conocido por una especie A. mirus representada por una única mandibular (p2-m2) procedente de las colecciones antiguas del Quercy, cuya edad geológica es des- conocida. Nuevo material procedente de la localidad de Pech-du-Fraysse completa nuestro conoci- miento sobre el género, en particular para el maxilar y la dentición superior. Adelpharctos pertenece a la subfamilia Hemicyoninae, familia Ursidae. El género difiere de los hemicioninos del Mioceno medio, que tienen dentición más robusta, y de los del grupo Cephalogae-Phoberogale por algunos caracteres morfológicos. Se interpreta como perteneciente a una línea procedente del grupo ancestral primitivo de la subfamilia Hemicyoninae. Palabras clave: Oligoceno superior, Francia, Quercy, Carnivora, Ursidae, Hemicyonidae.

Introduction museums through the whole world. The sediments and the fossils were provided by many quarries dis- Although they were used from the last third of persed over a large surface of hundreds of km2. the 19th century to the beginning of the 20th to Several palaeontological studies of different mam- extract phosphate used as fertilizer for agricultural mals (Gervais 1872 a & b; Filhol 1872 a & b, 1874, needs, the fissure, cave and chasm fillings of the 1876, 1877, 1880, 1882; Gaillard 1908; Schlosser “Phosphorites du Quercy” are essentially well 1888, 1889, 1891) leaded to the conclusion that the known because the myriads of fossils which were deposits may represent a large range of time from mixed with the sediments. Among these fossils, the late Eocene to late Oligocene but that, except in thousands of vertebrate bones, particularly mammal rare cases, the fossils were mixed in every locality. bones, were unearthed and stored in number of Other studies especially focused on Carnivora of

1 IPHEP, UMR 6046 du CNRS, UFR Sciences, Université de Poitiers, 40 Avenue du recteur Pineau, 86022 Poitiers, France. Email: [email protected] e390-11 De Bonis.qxd 30/1/12 14:15 Página 180

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the old collections (Teilhard 1915; Piveteau 1931, Material and methods 1942/1943, 1962; Bonis 1966, 1971, 1981; Gins- burg 1966, 1979; Lange-Badré 1969, 1970; Peigné The studied material comes from the locality of Pech-du- 2000, 2001, 2003; Peigné & Bonis 1999; Wolsan & Fraysse (Saint Projet, Tarn-et-Garonne, France) in the military camp of Caylus (Crochet 1971). It comprises a right hemi- Lange-Badré 1996) could not clear the problem of mandible with the alveolus of i3 and c, p1, broken p2, p3-m2, the fossil dating as well. In fact, the names some- alveolus of m3 and a right maxilla with an alveolus of P1 and times labelled on the fossils (Mouillac, Lamandine, P2-M2. The two specimens have been found close together Saint Antonin, Prajoux, Mémerlin…) correspond to with a similar wear pattern and they clearly belong to the same quite large areas containing many fillings whose individual. The fauna contains many species of small and large vertebrates which allow a precise biochronological dating and geological ages may be very different. Thus the thus the locality is the reference biochronological level MP 28 material of the old collections which was listed in a (see Remy et al. Biochrom 97 p. 792-793) about 25-26 Ma old. catalogue (Sigé et al. 1979) was a taxonomic melt- The order Carnivora is well represented in the old collections ing pot without any indication of the chronology as well as in the new ones (Bonis 1974, 1976, 1978; Bonis & despite some very complete and nice specimens. Cirot 1995; Cirot & Bonis 1992, 1993; Peigné & Bonis 1999, 2003) and the genus Adelpharctos as well as the type-species A. Moreover, many people thank that the works were mirus were described from a well preserved but unique stopped in the Quercy because all the fissures were mandible of the old collections (Bonis 1971). The new material empty without any phosphorite or fossil. completes our knowledge on the anatomy and biostratigraphic Nevertheless, a new phase of researches began position of this genus. The measurements were taken with an before the second world war, when Gèze (1938 a, electronic calliper to 1/100° and given to 1/10°. 1938 b) demonstrated that each of two fissure fillings quite close contained a homogeneous fauna. During the second half of the sixties, a scientific Systematics team coming from the universities of Montpellier II (USTL), Paris 6 (UPMC), Lyon 1 (Claude Order Carnivora Bodwich, 1821 Family Ursidae Fischer de Waldheim, 1817 Bernard), the Muséum national d’Histoire naturelle Sub-family Hemicyoninae Frick, 1926 Paris and later the University of Poitiers carried out Genus: Adelpharctos Bonis, 1971 several field campaigns in the Quercy which gave Type species: Adelpharctos mirus Bonis, 1971 rise to a large number of general publications on the Other species: A. ginsburgi n.sp. Quercy phosphorites faunas (Bonis et al. 1973; Bonis et al. 1974; Bonis et al. 1977; Crochet et al. Emended diagnosis: Dental formula: 3/3 I, 1/1 C, 4/4 P, 2/3 M. Hemicyoninae with a more cutting dentition than the other 1975; Crochet et al. 1981; Hartenberger et al. 1974; genera of the sub-family, m1 shorter relative to the premolar Gèze et al. 1978; Remy et al. 1987) and many other row with a less reduced metaconid and a talonid tapering back- ones showed that every fissure had an original ward. P2-3 with three roots, P4 short and robust, triangular fauna and of which two of them may precise the shaped M1 with a metaconule more buccal than the protocone, a hypometacrista joining the base of the metacone and with a global geological age of the Phosphorites du Quer- modest linguo-distal cingulum. cy which are now dated from the middle Eocene (Astruc et al. 2000) to the early Miocene (Sigé et Adelpharctos ginsburgi n. sp. al. 1991). In every site, the fossiliferous levels are Holotype: right mandible with alveolus of i3, c-m2, alveolus quite shallow and deposited during a relatively of m3 and right maxilla with alveolus of C-P1 and P2-M2 short time. The amount of variation in every species (PFRA 34). Locality: Pech-du-Fraysse, Lot (France) is similar to that of extant species (Vianey-Liaud & Dating: Late Oligocene (MP 28) Legendre 1986). Because the large number of sites Origin of the name: dedicated to my colleague the late and the possibilities of evolutionary studies, the Léonard Ginsburg for his numerous works on the carnivorans. Quercy phosphorites constitute a true open air labo- Diagnosis: Adelpharctos smaller than the type-species A. ratory of vertebrate evolution (Bonis et al. 1977). mirus, p4 slightly less robust with a minute posterior accessory cuspid, metaconid of m1 slightly less posterior, talonid basin of There are several fossil bearing localities which m1 slightly less reduced. are used as level markers in the frame of the conti- nental Paleogene biochronology establishing a scale founded on the evolution of mammalian faunas as Description are the American “Ages of mammals”. Some of the mammalian level markers are Quercy phosphorites Mandible (Fig. 1-2; Table 2): The ramus is complete except localities (see Biochrom 1997). a small portion of the coronoid process. The corpus mandibu-

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A new species of Adelpharctos (Mammalia, Carnivora, Ursidae) 181

Table 1.—l = length; w = width; trL = trigonid length; trw = trigonid width; talw = talonid width

Lower teeth cL cw p1L p1w p2L p2w p3L p3w p4L p4w m1L m1trL m1trw m1talw m2L m2trw m2talw

A. mirus 9,9 4,6 12,3 6,8 19,3 14,2 8 7,7 10,8 7,2 6,3 A. ginsburgi 9,3 6,1 5,7 3,9 9,8 3,7 10,5 5,4 17,2 12,7 7,5 7,3 9,9 6,5 5,4

Fig. 1.—Adelpharctos ginsburgi n. sp. Right mandible (PFRA 35); A: buccal view, B: lingual view (Scale bar = 3 cm).

lae is slightly elongate with a quite short symphysis; there are two small mental foramina, one anterior and situated beneath the mesial root of p2, another one posterior beneath the mesial root of p3, both at midheight of the ramus. In the lingual surface, the foramen mandibulare is relatively small and opens 21 mm in front of the condyle. The ventral border is slightly convex under the tooth raw, the deeper part being under m2, and it turns upward under the coronoid process. The latter is high with a deep masseteric fossa limited downward by a rough crest and extending to the alveolus of m3. The condyle is slightly oblique relative to the coronoid process and level with the tooth row. The angular process, long and robust is back- ward and downward directed; its dorsal surface is relatively wide and its posterior-most part turns upward like a hook. Lower teeth (Fig. 1; Table 1): The incisors are lost and the remaining i3 alveolus is small and oval shaped with a vertical great axis. The canine is relatively thin and long. The buccal surface is convex but the lingual one is partially concave with, at the mesio-lingual corner, a faint ridge which extends from the tip to the basis and runs to the distal end. There is a 1 mm diastema from the canine to p1 which is one-rooted, small, elongate and narrow with an anteriorly situated main cuspid. Fig. 2.—Adelpharctos ginsburgi n. sp. Right mandible (PFRA The other premolars have two roots; p2 is partially broken off, 35); occlusal view (Scale bar = 1 cm).

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Table 2.—Height md = height of the mandible beneath

Height md p1-p2 p2-p3 p3-p4 p4-m1 m1-m2

A. mirus 19,1 19,3 20,6 22,5 A. ginsburgi 16,8 17,7 18,9 20,6 23,1

Fig. 3.—Adelpharctos ginsburgi n. sp. Right maxilla (PFRA 36); A: buccal view, B: lingual view (Scale bar = 1 cm).

while p3 and p4 are complete; both are higher than the paraconid of m1, especially p4; the latter has a small but distinct pacd at midheight. The carnassial is moderately elongate; the paraconid is low, short and oblique relative to the tooth row; the protoconid is tall and relatively short; the lingual surface of the protoconid is a vertical semi-pillar with two vertical mesial and distal strips; the metaconid, quite well developed, is clearly separated from the paraconid and slightly posteriorly situated; the talonid is short for a hemicyonine with a high, quite large and sub-conical hypoconid and a low crest surrounding the talonid basin until the distal basis of the metaconid. The trigonid of m2 is essentially composed by the protoconid and metaconid which have quite the same volume and are situated at the same level; a small anterior bump corresponds to a vestigial paraconid; the protoconid is separated by a notch from the hypoconid while a entoconid ridge runs without notch from the metaconid an joins the mesial face of the hypoconid, the talonid basin being very small and narrow. An oval alveolus indicates a slightly elongate small one-rooted m3. Maxilla (Fig. 3): The premaxilla is absent and only the lower part of the maxilla is preserved. A large infraorbital foramen is situated just above the top of the mesio-buccal root of P4, at Fig. 4.—Adelpharctos ginsburgi n. sp. Right maxilla (PFRA 36); 10.8 mm of the lingual margin. occlusal view (Scale bar = 1 cm).

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A new species of Adelpharctos (Mammalia, Carnivora, Ursidae) 183

Table 3.—pcl = paracone length; msl = metastyle length; msw = metastyle width; mw = mesial width; dw = distal width

Upper teeeth P2L P2w P3L P3w P4L P4w P4pcL P4msL P4msw M1L M1mew M1dsw M2L M2w

A. ginsburgi 8,2 5,5 10,5 5,9 15,4 11,4 6,6 6 5,8 11,9 16,7 14,9 7,3 12,3

Upper teeth (Fig. 3-4; Table 3): There a large but not very pacd and particularly the shape of m2 with a metaconid as devel- deep alveolus for the root of the canine; the maxilla is preserved oped as or more developed than the protoconid. Adelpharctos is on 9.4 mm in front of this alveolus without any trace of an nevertheless clearly distinct from the Amphicynodontidae by the incisor alveolus; thus the diastema was larger. There is only a longer m1 relative to the whole tooth row (see Teilhard 1915, p. 1.5 mm diastema between the canine and the alveolus of a one- 50). Most Ursinae have less tall premolars and more elongate rooted P1. P2 is separated from P1 by less than 3 mm; it has a molars than those of Adelpharctos. The oldest ursine genus Bal- simple asymmetric crown, the mesial part being smaller than the lusia (MN 3, Lower Miocene) differs from Adelpharctos by, in distal one, without any posterior accessory cusp (pac), but with addition to the double rooted upper P2 and P3, the less tall and a small anterior bulging at the basis of the crown and a slightly less trenchant premolars, less tall m1 with a longer and flat larger distal one, and a thin crest running in the mesial and distal talonid, shorter P4 and more elongate sub-quadrate M1 with a edges of the crown; there are three roots lined mesio-distally, metaconule level with the protocone (Ginsburg & Morales the central one being shorter than the others. P3 is separated 1998). The Pech-du-Fraysse carnivore shows more similarities from P2 by a 5.4 mm diastema: the shape of the crown is similar with the Hemicyoninae whose dentition is less hypocarnivore to that of P2 but it is less asymmetrical with more developed than that of the Ursinae. Zaragocyon Ginsburg & Morales, 1995 bulging at the base and it is as tall as the paracone of P4; it has is slightly larger (m1 = 19.8 mm) and differs from Adelpharctos also three roots but the central one is more lingually situated, by the upper carnassial without parastyle, shorter relative to the corresponding to a lingual bulging of the crown. The short P4 is molars, with a more distally situated protocone, the elongated separated from P3 by 2 mm; there is a moderate and bulbous and sub-quadrate M1 whose metaconule is level to the paracone, parastyle; the sub-conical paracone is asymmetrical, the mesial the more developed M2 with a larger lingual cingulum; the low part being longer than the distal one; it is followed by a short lower premolars, low protoconid, inflated and distally situated metastyle blade; the protocone is well developed but without metaconid and larger talonid of m1; the relatively longer m2. cusp, composed by a hollow basin surrounded by a blunt cingu- Phoberocyon differs by the larger size (m1 = 24.6 to 35 mm. lar ridge which continues along the metastyle blade; a faint cin- depending on the species), the more distally situated protocone gulum runs along the buccal side. M1 is triangular, the meta- and the lack of parastyle of P4, the larger M1 and M2 with cone is more lingually situated and smaller than the paracone larger lingual cingulum, the less tall lower p3 and p4 with a and the mesial face is wider than the distal one, thus the buccal more developed pacd, the lower protoconid, reduced metaconid face is oblique; the protocone, small and low, is far from the lin- and wider talonid of m1, the longer m2. The close genus Hemi- gual corner; the protocrista is short and vanishes when contacts cyon differs from Adelpharctos by quite the same characters, as the paracone, the metacrista is thin but slightly longer with a do Plithocyon and Dinocyon. small but well distinct metaconule situated more buccally than Other Hemicyoninae fit better the Pech-du-Fraysse carnivore the protocone; there is no mesial cingulum but a swelling in the and have been attributed to the with genera Cephalogale Jour- disto-lingual corner; a faint cingulum is present along the buccal dan, 1862 or Phoberogale Ginsburg & Morales, 1995. Represen- surface. M2 is smaller than M1 and less asymmetrical; it is oval tatives of this group are known from the early Oligocene (MP shaped, the buccal length being slightly smaller than the lingual 21) to the early Miocene (MN 1-2). The geological oldest species one; the paracone is the larger buccal cusp, the métacone being are quite small (m1 = about 12 mm.) but the size increases minute, both are slow and quite fused; the protocone is small through the time and the younger species are middle sized hemi- with two delicate trigone crests in which it is impossible to dis- cyonines (m1 = 21 to 23 mm.) with larger m1 relative to the pre- tinguish any conule; both crests vanish at the base of the buccal carnassial premolars and longer m2. All these species differ by cusps; there is a moderate cingulum surrounding the protocone less tall pre-carnassial premolars, absence of parastyle in P4, along the lingual surface and a very faint along the buccal one. more sub-quadrate M1 whose lingual cingulum is more anteriorly developed, absence of hypometacrista, postprotocone crista joining directly the distal cingulum with or without distinct meta- Comparisons conule, and generally more reduced m1 metaconid. Nevertheless Adelpharctos, by its overall morphology, is closer to Cephalo- gale than to another genus and belongs to a group different from There are some small differences between Adelpharctos gins- the other hemicyonines which appeared in Europe in the limit burgi and A. mirus, thus the comparisons are made from the early-middle Miocene (MN 3) with a more massive dentition. genus Adelpharctos as a whole. The Amphicyonidae essentially differ from Adelpharctos by the presence of M3, the double-rooted upper premolars P2 and P3, the longer upper carnassial with a reduced protocone, the more trenchant m1 and more developed Conclusions protoconid of m2. The overall morphology of Adelpharctos leads to compare to the Ursidae. Adelpharctos shows some features found in the Amphicynodontidae, the probably polyphyletic stem The new material of Pech-du-Fraysse allows the group in which the Ursidae are rooted, like the lack of developed description of a new species of a primitive ursid,

Estudios Geológicos, 67(2), 179-186, julio-diciembre 2011. ISSN: 0367-0449. doi:10.3989/egeol.40553.181 e390-11 De Bonis.qxd 30/1/12 14:15 Página 184

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Adelpharctos ginsburgi n. sp.. It is smaller than A. these taxa may be considered as belonging to the mirus, type species of the genus and different by bulk of the whole oligocene local fauna. But Amphic- some characters (p4 slightly relatively less robust tis and Stenoplesictis crocheti seem to be new immi- with a minute posterior accessory cuspid, metaconid grants in the European late Oligocene as is Adel- of m1 slightly less posterior, talonid basin of m1 pharctos. They rapidly were followed (MP 29) by the slightly less reduced). Closer relationships do exist amphicyonids Haplocyon and Haplocyonopsis as with the genera Cephalogale and Phoberogale. The well as the small anthracotheriid Microbunodon). All size was increasing in several lineages of Cephalo- these taxa may belong to “the late travellers of the gale during the Oligocene and early Miocene (Bonis big Stampian migration” (Viret 1929) which changed 1973). If the same did occur in the genus Adelpharc- the core of the European fauna after the “Grande tos, A. mirus would be geologically younger than A. Coupure” of Stehlin (1909). We don’t know the roots ginsburgi n.sp. of this late Oligocene migration but its origin is prob- If we look for the place of Adelpharctos, Cephalo- ably in Asia although we don’t know any direct gale and Phoberogale in the guild of Oligocene car- ancestor in this area for many of the taxa. nivorans, we see that their overall dental characteris- tics are quite similar to those of extant Canidae with quite large lower carnassials and close proportions ACKNOWLEDGEMENTS between trigonid and talonid. Their size, based on The new material from the Quercy phosphorites was m1 length, vary from those of small foxes like obtained by the works of a team sponsored by the French Vulpes rueppeli or V. corsac to that of the coyote, CNRS (Recherches Cooperatives sur Programme) and helped Canis latrans. As far as we know the limb skeleton by several French universities. I wish to thank the military of this group, “It is characterized by its cursorial authorities which allowed our team to work in the military adaptations, such as digitigrade posture…” (Wang & camp of Caylus. I am indebted to many colleagues, technicians and students who took part in these works but I would like to al. 2009), thus the Adelpharctos-Cephalogale- cite especially my colleague Jean-Yves Crochet for numbers of Phoberogale group probably occupied during the field campaigns in the locality of Pech-du-Fraysse. I thank also Oligocene and part of the early Miocene some eco- the members of the IPHEP (University of Poitiers) in which logical niches close to those of extant canids, foxes, this article was prepared and J. Morales to invite me to present jackals or coyotes. The high number of Cephalogale this paper in honour to our colleague L. Ginsburg. found in some quarries could indicate that they were living in packs of several individuals. Other contes- tant carnivorans could have been small amphicy- References onids (Cynelos or Goupilictis) which were less abundant and fully plantigrades or the primitive felid Astruc, J.G.; Augé, M.; Bonis, L. de; Brunet, M.; Cirot, Proailurus whose way of hunting certainly was very E.; Compte B.; Crochet, J.Y.; Lange-Badré, B.; different. The large amphicyonids were certainly Legendre, S.; Lévèque F.; Marandat, B.; Mourer- Chauviré, C.; Rage, J.C.; Remy, J.A.; Sigé, B.; Sudre, feeding on larger preys. The favourite preys of this J. & Vianey-Liaud M. (1995). Le Garouillas et les group could have been large theridomyid rodents, sites contemporains (Oligocène, MP 25) des phospho- cainotheriids and primitive cervoids of the sheep- rites du Quercy (Lot, Tarn-et-Garonne, France) et sized genera Dremotherium and Bedenomeryx which leurs faunes de vertébrés. Palaeontographica Stuttgart, are generally very abundant in the fossiliferous A 236, 343 pp. localities. We may notice that the disappearance of Astruc, J.-G.; Escarguel, G.; Marandat, B.; Simon- this group of carnivorans corresponds to the disap- Coinçon, R. & Sigé, B. (2000). Floor-age constrai- ning of a tectonic paroxism of the Pyrenean orogen. pearance or rarefaction of these preys. Late middle Eocene mammal age of a faulted karstic The fauna from the locality of Pech-du-Fraysse filling of the Quercy phosphorites, south-western contains until now 8 species of Carnivora. Adel- France, Geodinamica Acta Paris, 13: 271-280. pharctos ginsburgi, Cephalogale sp., Amphictis doi:10.1016/S0985-3111(00)01047-0 ambigua, Amphictis cf. nana, “Plesictis” stenogali- Biochrom 97 (1997). Synthèses et tableaux de corréla- nus, Amphicyonidae sp., Stenoplesictis cayluxi, tions. In: Actes du Congrès Biochrom 97, (J.P. Aguilar, S. Legendre, & J. Michaud, eds.). Mémoires et Tra- Stenoplesictis crocheti. Amphicyonidae are common vaux de l’EPHE, Montpellier, 21, 767-805. in the European Oligocene, Cephalogale was present Bonis, L. de (1966).Arrières-Crânes et moulages from the early Oligocene (Mas de Got MP 22) as was endocrâniens de Carnivores fossiles. Annales de Palé- Stenoplesictis cayluxi (Itardies MP 23), and thus ontologie Paris, 52 (2): 143-162.

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Bonis, L. de (1971). Deux nouveaux carnassiers des Coupure” découvertes dans les phosphorites du Phosphorites du Quercy. Annales de Paléontologie Quercy. Bulletin du Muséum national d’Histoire natu- Paris, 57 (1): 117-127. relle Paris, 3 (C): 245-266. Bonis, L. de (1973). Contribution à l’étude des mam- Delfortrie, M. (1872). Les gîtes de chaux phosphatée mifères de l’Aquitanien de l’Agenais. Rongeurs, Péris- dans le département du Lot, leur faune, le mode et l’é- sodactyles, Carnivores. Mémoires du Muséum national poque probable de leur formation. Actes de la Société d’Histoire naturelle Paris, Sciences de la Terre, n.s. C, Linnéenne de Bordeaux Bordeaux: 503-518. 26: 1-192. Filhol, H. (1872 a). Sur les carnassiers et les chiroptères Bonis, L. de (1974). Premières données sur les carnivo- dont on trouve les débris fossiles dans les gisements res fissipèdes provenant des fouilles récentes dans le des phosphorites de Caylux, Fregols et Concots. Jour- Quercy. Palaeovertebrata Montpellier, 6: 27-32. nal de Zoologie Paris, 1: 280-283. Bonis, L. de (1976). Découverte d’un crâne d’Amphictis Filhol, H. (1872 b). Recherches sur les mammifères fos- (Mammalia, Carnivora) dans l’Oligocène supérieur des siles des dépôts de phosphate de chaux dans les dépar- Phosphorites du Quercy. Comptes rendus de l’Acadé- tements du Lot, du Tarn et de Tarn-et-Garonne. Anna- mie des Sciences Paris, 283(4): 327-330. les des Sciences Géologiques Paris, 3: 1-31. Bonis, L. de (1978). La poche à phosphate de Ste Nébou- Filhol, H. (1874). Nouvelles observations sur les mam- le (Lot) et sa faune de vertébrés du Ludien supérieur. mifères des gisements de phosphates de chaux, Lému- 12. Fissipèdes (Carnivores). Palaeovertebrata Montpe- riens et Pachylémuriens. Annales des Sciences Géolo- llier, 8 (2/4): 301-311. giques Paris, 4: 1-36. Bonis, L. de (1981). Contribution à l’étude du genre Filhol, H. (1876). Recherches sur les Phosphorites du Palaeogale Meyer (Mammalia, Carnivora). Annales de Quercy. Etude des fossiles qu’on y rencontre et spécia- Paléontologie Paris, 67 (1): 37-56. lement des mammifères. Première partie, Bibliothèque Bonis, L. de; Brunet, M.; Cavaillé, A.; Crochet, J.Y.; Gèze, de l’Ecole des Hautes Etudes, Sciences Naturelles B.; Ginsburg, L.; Lopez, N.; Hartenberger, J.L.; Rage, Paris, 15 (4) :1-220. J.C.; Remy, J.A.; Sigé, B.; Sudre, J.; Thaler, L. & Via- Filhol, H. (1877). Recherches sur les Phosphorites du ney-Liaud, M. (1974). Table ronde sur les Phosphorites Quercy. Etude des fossiles qu’on y rencontre et spécia- du Quercy. Palaeovertebrata Monpellier, 6 (1-4): 1-303. lement des mammifères. Seconde partie, Bibliothèque Bonis, L. de & Cirot, E. (1995). Le Garouillas et les sites de l’Ecole des Hautes Etudes, Sciences Naturelles contemporains (Oligocène, MP 25) des Phosphorites Paris, 16 (1): 1-338. du Quercy (Lot, Tarn-et-Garonne, France) et leurs fau- Filhol, H. (1880). Découverte de mammifères nouveaux nes de Vertébrés. 7. Carnivores. Palaeontographica dans les dépôts de phosphate de chaux du Quercy. Stuttgart, 236 (1-6): 135-149. Comptes Rendus de l’Académie des Sciences Paris, 91: Bonis, L. de; Crochet, J.Y.; Rage, J.C.; Sigé, B.; Sudre, 344-346. J. & Vianey-Liaud, M. (1973). Nouvelles faunes de Filhol, H. (1882). Mémoires sur quelques mammifères vertébrés oligocènes des phosphorites du Quercy. fossiles des phosphorites du Quercy. Vialelle Toulou- Bulletin du Muséum national d’Histoire naturelle se, 140 pp. Paris, (3) 174 (Sciences de la Terre 28): 103-113. Filhol, H. (1894). Observations concernant quelques Bonis, L. de; Crochet, J.Y.; Hartenberger, J.L.; Rage, J.C.; mammifères fossiles nouveaux du Quercy. Annales des Sigé, B.; Sudre, J. & Vianey-Liaud, M. (1977). Les Sciences Naturelles: Zoologie et Paléontologie Paris, Phosphorites du Quercy: un laboratoire naturel de l’évo- 16: 129-150. lution des vertébrés. Courrier du CNRS Paris, 25: 6-11. Frick, C. 1926. The Hemicyoninae and an American Ter- Bowdich, T.E. 1821. An analysis of the Natural Classifi- tiary bear. Bulletin of the American Museum of Natural cation of Mammals for the use of Students and trave- History, 56: 1-119. llers. 115 ppp. J. Smith, Paris. Gervais, P. (1872 a). Sur les mammifères dont les osse- Cirot, E. & Bonis, L. de (1992). Revision du genre Amp- ments accompagnent les dépôts de chaux phosphatée hicynodon, carnivore de l’Oligocène. Palaeontograp- du Tarn-et-Garonne et du Lot. Comptes Rendus de l’A- hica Stuttgart, 220(4-6): 103-130. cadémie des Sciences Paris, 74: 1367-1371. Cirot, E. & Bonis, L. de (1993). Le Crâne d’Amphictis Fischer von Waldheim, G. 1813. Zoognosia tabulis ambiguus (Carnivora, Mammalia): son importance synopticis illustrata. Moscow, Nicolai SErgeidis Vse- pour la compréhension de la phylogénie des Mustéloï- volozsky, vol. 2, 605 pp. des. Comptes rendus de l’Académie des Sciences Paris, Gervais, P. (1872b). Sur les mammifères dont les osse- 316: 1327-1333. ments accompagnent les dépots de chaux phosphatée Crochet, J.Y. (1971). Les vertébrés de l’Oligocène supé- des départements de Tarn et Garonne et du Lot. Jour- rieur du Pech de Fraysse, poche à phosphate du Quercy nal de Zoologie Paris, 1: 261-268. (commune de Saint-Projet, Tarn-et-Garonne). Comptes Gèze, B. (1938a). Contribution à la connaissance des Rendus sommaires des Séances de la Société Géologi- Phosphorites du Quercy. Bulletin de la Société Géolo- que de France Paris, 9: 316-317. gique de France Paris, 5 (8): 123-146. Crochet, J.Y.; Hartenberger, J.L.; Rage, J.C.; Remy, J.A.; Gèze, B. (1938b). La genèse des Phosphorites du Sigé, B.; Sudre, J. & Vianey-Liaud, M. (1981). Les Quercy. Comptes Rendus de l’Académie des Sciences nouvelles faunes de vertébrés antérieures à la “Grande Paris, 206: 759-761.

Estudios Geológicos, 67(2), 179-186, julio-diciembre 2011. ISSN: 0367-0449. doi:10.3989/egeol.40553.181 e390-11 De Bonis.qxd 30/1/12 14:15 Página 186

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Gèze, B.; Rage, J.C.; Vergnaud-Grazzini, C.; Broin, F. relations avec N. brachyops d’Amérique du Nord. de; Buffetaud, E.; Moure-Chauviré, C.; Crochet, J.Y.; Revue de Paléobiologie, 18 (1): 57-67. Sigé, B.; Sude, J.; Remy, J.A.; Lange-Badré, B.; Bonis, Peigné, S. & Bonis, L. de (1999 b). The genus Stenople- L. de; Hartenberger, J.L. & Vianey-Liaud, M. (1978). sictis Filhol (Mammalia, Carnivora) from the Oligoce- La poche à phosphate de Sainte-Néboule (Lot) et sa ne deposits of the Phosphorites du Quercy, France. faune de vertébrés du Ludien supérieur, Palaeoverte- Journal of Vertebrate Paleontology, 19 (3): 566-575. brata Montpellier 8 (2-4): 167-326. doi:10.1080/02724634.1999.10011165 Gaillard, C. (1908). Les oiseaux des phosphorites du Peigné, S. & Bonis, L. de (2003). Juvenile cranial ana- Quercy. Annales de l’Université de Lyon, 1 (23): 1-178. tomy of Nimravidae (Mammalia, Carnivora): biologi- Ginsburg, L. (1966). Les Amphicyons des Phosphorites cal and phylogenetic implications. Zoological Journal du Quercy. Annales de Paléontologie (Vert.) Paris, 52 of the Linnean Society London, 138: 477-493. (1): 23-44. doi:10.1046/j.1096-3642.2003.00066.x Ginsburg, L. (1979). Révision taxonomique des Nimravi- Piveteau, J. (1931). Les chats des Phosphorites du ni (Carnivora, Felidae) de l’Oligocène des Phosphori- Quercy. Annales de Paléontologie, 20: 107-163. tes du Quercy. Bulletin du Muséum national d’Histoire Piveteau, J. (1942/1943). Etudes sur quelques mammifè- naturelle Paris, 1 (1): 35-49. res des Phosphorites du Quercy. I. Le genre Stenople- Ginsburg, L. & Morales, J. (1995). Zaragocyon daamsi sictis. Annales de Paléontologie, 30: 63-72. n. gen. sp. nov., Ursidae primitif du Miocène inférieur Piveteau, J. (1962). L’encéphale de Viverravus angusti- d’Espagne. Comptes rendus de l’Académie des Scien- dens, Miacidé des Phosphorites du Quercy. Annales de ces Paris, 321 série IIa: 811-815. doi:10.1016/S0753- Paléontologie, 48: 165-175. 3969(98)80003-7 Remy, J.A.; Crochet, J.Y.; Sigé, B., Sudre, J.; Bonis, L. Ginsburg, L. & Morales, J. (1998). Les Hemicyoninae de; Vianey-Liaud, M.; Godinot, M.; Hartenberger, J. (Ursidae, Carnivora, Mammalia) et les formes appa- L.; Lange-Badré, B.& Comte, B. (1987). Biochronolo- rentées du Miocène inférieur et moyen d’Europe occi- gie des Phosphorites du Quercy: Mise à jour des listes dentale. Annales de Paléontologie, 84 (1): 71-125. faunistiques et nouveaux gisements de mammifères Hartenberger, J.L.; Sigé, B. & Sudre, J. (1974). La plus fossiles. Münchner Geowissenschaftliche Abhandlun- ancienne faune de mammifères du Quercy: Le Bretou. gen, 10: 169-188. Palaeovertebrata, 6 (3-4):177-196. Schlosser, M. (1888). Die Affen, Lemuren, Chiropteren, Lange, B. (1969). Un nouveau musteliné des Phosphori- Insectivoren, Marsupialier, Creodonten und Carnivo- tes du Quercy, Mustelictis piveteaui. Comptes rendus ren des europäischen Tertiärs und deren Beziehungen de l’Académie des Sciences Paris, 268: 2870-2872. zu ihren lebenden und fossilen aussereuropäischen Lange, B. (1970). Mustelictis piveteaui, Mustélidé nou- Verwandten. Beiträge zur Paläontologie Österreich veau des Phosphorites du Quercy. Annales de Paléon- Ungarns und des Orients, I. Theil 6: 1-224. (1889). II tologie, 56 (1): 3-16. Theil 7: 1-162. (1891). III Theil 8: 1-102. Legendre, S.; Sigé, B.; Astruc, J.-G.; Bonis, L. de; Cro- Sigé, B.; Aguilar, J.P. & Marandat, B. (1991). Extension chet, J.-Y.; Denys, C.; Godinot, M.; Hartenberger, J.- au Miocène inférieur des remplissages phosphatés du L.; Lévèque, F.; Marandat, B.; Mourer-Chauvire, C.; Quercy, la faune de vertébrés de Crémat (Lot, France). Rage, J.C.; Remy, J.A.; Sudre, J. & Vianey-Liaud, M. Geobios, 23 (4): 497-502. (1997). Les phosphorites du Quercy: 30 ans de recher- Stehlin, H.G. (1909). Remarques sur les faunules de che. Bilan et perspectives In: Actualités paléontologi- mammifères des couches éocènes et oligocènes du ques en l’honneur de Claude Babin (Racheboeuf P.R. Bassin de Paris. Bulletin de la Société Géologique de & Gayet M., Eds), Géobios 20, 331-345. France, 9 (4): 488-520. Peigné, S. (2000). A new species of Eofelis (Carnivora: Vianey-Liaud, M. & Legendre, S. (1986). Les faunes des Nimravidae) from the Phosphorites of Quercy, France. phosphorites du Quercy: principes méthodologiques en Comptes Rendus de l’Académie des Sciences Paris, paléontologie des mammifères; homogénéité chronolo- 330 (9): 653-658. gique des gisements de mammifères fossiles, Eclogae Peigné S. (2001). A primitive nimravine skull from the geologicae Helvetiae, 79 (3): 917-944. Quercy fissures, France. Implications for the origins Wang, X.; Hunt, R.M.; Tedford, R.H. & Lander, E.B. and evolution of Nimravidae (Carnivora). Zoological (2009). First record of immigrant Phoberogale (Mamma- Journal of the Linnean Society London, 132: 401-410. lia, Ursidae, Carnivora) from Southern California. Geodi- doi:10.1111/j.1096-3642.2001.tb02467.x versitas 31 (4): 753-773. doi:10.5252/ g2009n4a753 Peigné, S. (2003). Systematic review of European Nim- Wolsan, M. & Lange-Badré, B. (1996). An arctomorph ravinae (Mammalia, Carnivora, Ninravidae) and the carnivoran skull from the Phosphorites du Quercy and phylogenetic relationships of Palaeogene Nimravidae. the origin of procyonids. Acta Palaeontologica Poloni- Zoologica Scripta, 32: 199-229. doi:10.1046/j.1463- ca, 41(3): 277-298. 6409.2003.00116.x Peigné, S. & Bonis, L. de (1999 a). Le premier crâne de Recibido el 10 de enero de 2011 Nimravus (Mammalia, Carnivora) d’Eurasie et ses Aceptado el 16 de mayo de 2011

Estudios Geológicos, 67(2), 179-186, julio-diciembre 2011. ISSN: 0367-0449. doi:10.3989/egeol.40553.181